Canopy and leaf gas exchange of <Emphasis Type="Italic">Haloxylon ammodendron</Emphasis> under different soil moisture regimes

In order to reveal the drought resistance and adaptation of the C₄ desert plant Haloxylon ammodendron under artificially controlled soil moisture regimes, representative plants were selected to measure canopy photosynthesis using canopy photosynthetic measurement system. The results showed that appropriate soil moisture significantly enhances the canopy and leaf photosynthetic capacity, and extremely high soil moisture is not conducive to the photosynthesis of H. ammodendron.     

固沙树种梭梭在不同水分梯度下的光合生理特征

对不同水分梯度下梭梭光合作用的研究发现,地下水位、土壤含水量、灌水量对梭梭的光合具有明显的影响。在1．4m、2．4m、3．4m地下水位下,梭梭的光合速率、呼吸作用、量子效率随水位的降低而降低,光补偿点随水位的降低而增大;在滴灌供水时,梭梭的光合速率较对照明显提高,且灌水量越大,梭梭的光合速率越高;半流动沙丘、丘间低地、粘土沙障固定沙丘退化梭梭的光合作用的强弱同其林地土壤水分的高低基本一致。7月,不同水分梯度下的梭梭光合作用均表现出午休现象,9月只有立地条件较差的梭梭发生光合午休,而且9月光合作用要强于7月,说明9月环境条件更有利于梭梭生长。     

民勤绿洲人工梭梭林退化现状、特征与恢复对策

民勤全县共有人工梭梭林近3．5万hm^2,占人工林总面积的51．5％,在民勤固沙林中占主导地位。但目前除水库与环河周围外,人工梭梭林均出现不同程度的退化现象,并表现为建群种梭梭光合强度减弱、群落盖度降低、梭梭种群优势度下降、林地土壤含水率下降、组成群落的物种多样性减少、地表土壤结皮破坏、风蚀作用增强等特征。梭梭林的退化,直接威胁到民勤绿洲的生存与发展,其恢复与重建成为当前一项十分重要而紧迫的任务,为此提出控制梭梭林密度、人工集雨、布设粘土或新型材料沙障、引进与培育抗旱新品种、特殊区域应用喷灌技术补水、补充水源不断恢复民勤地区的地下水位等措施,以期达到恢复与维持梭梭群落稳定发展的目的。     

临泽绿洲边缘区棉花群体光合速率、蒸腾速率及水分利用效率

在甘肃河西走廊中部黑河中游绿洲边缘区,于6月下旬和8月上旬,利用Li-8100土壤碳通量测定系统与改进的同化箱联合对田间条件下早熟陆地棉(Gossypium hirsutum)品种新陆早8号的群体光合特性进行了研究.结果表明：试验地6月下旬的土壤呼吸速率和土壤蒸发速率显著高于8月上旬（P＜0.01）;棉花群体光合速率日变化均呈“单峰型”,6月下旬的群体光合速率显著高于8月上旬,其日平均值分别为（43.11±1.26）和（24.53±0.60）μmol CO₂·m^-2·s^-1, 差异极显著（P＜0.01）;群体蒸腾速率日变化也呈“单峰型”,6月下旬和8月上旬的日平均值分别为（3.10±0.34）和（1.60±0.26）mmol H₂O·m^-2·s^-1,两者存在极显著差异（P＜0.01）;6月下旬和8月上旬的群体水分利用效率日平均值分别为（15.67±1.77）和（23.08±5.54） mmol CO₂·mol^-1 H₂O,但差异不显著（P＞0.05）.两生育时期棉花群体光合速率与温度、光合有效辐射及土壤含水量均呈正相关关系.表明棉花群体光合速率在6月下旬和8月上旬均没有出现中午光合下调,8月由于土壤水分降低和植物叶片衰老,棉花群体光合速率和蒸腾速率显著降低,但水分利用效率并无显著下降.     

不同土壤水分条件下中国沙棘雌雄株光合作用、蒸腾作用及水分利用效率特征

采用LI-6400便携式光合系统对野外不同生境(沟底、坡面)和田间不同土壤水分条件下中国沙棘雌雄株的净光合速率和蒸腾速率及水分利用效率特征进行了观测,结果表明,中国沙棘雌雄株在水分条件较好的生境中均表现较强的生活力,净光合速率和蒸腾速率主要受光照强度和大气温度的影响,雄株表现出更高的光合、蒸腾、水分利用效率;在水分条件较差的生境中,雌雄株均通过降低蒸腾和提高水分利用效率来适应逆境,净光合速率和蒸腾速率的影响因子增多,雄株的光合速率大幅度下降,雌株仍保持较高的光合速率,雌株的水分利用效率高于雄株.可见,在反映植物瞬时生理变化的性状方面,雌株对土壤水分减少表现出了更强的适应性,雄株对土壤水分变化表现出了更大的敏感性.     

A model of dynamics of leaves and nitrogen in a plant canopy: an integration of canopy photosynthesis,leaf life span,and nitrogen use efficiency

A model of dynamics of leaves and nitrogen is developed to predict the effect of environmental and ecophysiological factors on the structure and photosynthesis of a plant canopy. In the model, leaf area in the canopy increases by the production of new leaves, which is proportional to the canopy photosynthetic rate, with canopy nitrogen increasing with uptake of nitrogen from soil. Then the optimal leaf area index (LAI; leaf area per ground area) that maximizes canopy photosynthesis is calculated. If leaf area is produced in excess, old leaves are eliminated with their nitrogen as dead leaves. Consequently, a new canopy having an optimal LAI and an optimal amount of nitrogen is obtained. Repeating these processes gives canopy growth. The model provides predictions of optimal LAI, canopy photosynthetic rates, leaf life span, nitrogen use efficiency, and also the responses of these factors to changes in nitrogen and light availability. Canopies are predicted to have a larger LAI and a higher canopy photosynthetic rate at a steady state under higher nutrient and/or light availabilities. Effects of species characteristics, such as photosynthetic nitrogen use efficiency and leaf mass per area, are also evaluated. The model predicts many empirically observed patterns for ecophysiological traits across species.     

古尔班通古特沙漠原生梭梭根区土壤水分变化特征

研究原生梭梭根区土壤水分动态变化对深入理解古尔班通古特沙漠水文过程与植被的相互作用具有重要意义.2007年8月至2008年7月,利用TDR土壤水分测定系统测定古尔班通古特沙漠南缘梭梭林地的土壤体积含水量,分析梭梭根区土壤水分时空分布及其变化规律.结果表明:梭梭根区土壤存在湿岛效应,根区60 cm土层水分储量是裸地的1.49倍,这种差异夏季比春季、降雨后比降雨前明显.春季积雪融化后,古尔班通古特沙漠土壤水分最丰富,而冬季的土壤水分含量最低.土壤水分年内变化可分为春季(3—5月)土壤水分迅速补充-消耗期、夏秋(6—9月)土壤水分缓慢消耗期和秋冬(10月至次年2月)土壤水分稳定期.经小波计算,根区和裸地土壤水分含量分别出现43 d、40 d的小周期和110 d、103 d的大周期.树冠的遮荫和自集水效应,以及根区土壤良好的渗透性是梭梭根区土壤水分相对丰富的主要原因.     

After more than a decade of soil moisture deficit,tropical rainforest trees maintain photosynthetic capacity,despite increased leaf respiration

Determining climate change feedbacks from tropical rainforests requires an understanding of how carbon gain through photosynthesis and loss through respiration will be altered. One of the key changes that tropical rainforests may experience under future climate change scenarios is reduced soil moisture availability. In this study we examine if and how both leaf photosynthesis and leaf dark respiration acclimate following more than 12 years of experimental soil moisture deficit, via a through‐fall exclusion experiment (TFE) in an eastern Amazonian rainforest. We find that experimentally drought‐stressed trees and taxa maintain the same maximum leaf photosynthetic capacity as trees in corresponding control forest, independent of their susceptibility to drought‐induced mortality. We hypothesize that photosynthetic capacity is maintained across all treatments and taxa to take advantage of short‐lived periods of high moisture availability, when stomatal conductance (g_s) and photosynthesis can increase rapidly, potentially compensating for reduced assimilate supply at other times. Average leaf dark respiration (R_d) was elevated in the TFE‐treated forest trees relative to the control by 28.2 ± 2.8% (mean ± one standard error). This mean R_d value was dominated by a 48.5 ± 3.6% increase in the R_d of drought‐sensitive taxa, and likely reflects the need for additional metabolic support required for stress‐related repair, and hydraulic or osmotic maintenance processes. Following soil moisture deficit that is maintained for several years, our data suggest that changes in respiration drive greater shifts in the canopy carbon balance, than changes in photosynthetic capacity.     

民勤沙区人工梭梭林自然稀疏过程研究

通过实地样方调查,对民勤沙区人工梭梭林的自然稀疏过程进行分析,以探讨其自然稀疏机理。结果表明:(1)民勤沙区梭梭经过自然稀疏,15~32 a林龄的密度为387.14株.hm-2。(2)梭梭林内乔木、灌木、草本植物对不同层次土壤水分利用的分异比较明显,草本植物和灌木分别主要利用0~20 cm和0~100 cm的土壤水分,梭梭主要利用60~120 cm的土壤水分。(3)在梭梭林成长过程中,沙丘水分递减,梭梭因水分亏缺而自然稀疏化,且在10 a林龄以内自然稀疏过程最明显。(4)当林龄达到15 a以上时,植株普遍枯梢,生长不良,植株个体增大缓慢。(5)对于15 a及其以上成龄梭梭林进行人工补灌没有显著恢复效果,只是增加草本植物的密度和盖度。     

矮嵩草草甸植物群落的光合特性研究

 矮嵩草草甸植物群落的光合、暗呼吸和土壤呼吸的研究表明：光合作用的日变化在6月份接近平坦型,7、8月份呈午间降低型。矮嵩草草甸植物群落的光合作用受较低的光合面积指数及冠层叶片的受光势态的影响,存在着明显的光饱和现象,光补偿点及光饱和点相对于全日照光合有效辐射均较低,接近于单叶的光响应特性。裸露地面的土壤呼吸和植物与土壤体系的暗呼吸不仅与温度有关,而且与土壤水分状况和降雨量也有密切联系。影响草甸群落光合特性的主要因素有：高原地区强烈的太阳辐射,较低的光合面积指数和植物根系与土壤紧密结合所形成的草结皮层结构。     

Intercropping Competition between Apple Trees and Crops in Agroforestry Systems on the Loess Plateau of China

Agroforestry has been widely practiced in the Loess Plateau region of China because of its prominent effects in reducing soil and water losses, improving land-use efficiency and increasing economic returns. However, the agroforestry practices may lead to competition between crops and trees for underground soil moisture and nutrients, and the trees on the canopy layer may also lead to shortage of light for crops. In order to minimize interspecific competition and maximize the benefits of tree-based intercropping systems, we studied photosynthesis, growth and yield of soybean (Glycine max L. Merr.) and peanut (Arachis hypogaea L.) by measuring photosynthetically active radiation, net photosynthetic rate, soil moisture and soil nutrients in a plantation of apple (Malus pumila M.) at a spacing of 4 m × 5 m on the Loess Plateau of China. The results showed that for both intercropping systems in the study region, soil moisture was the primary factor affecting the crop yields followed by light. Deficiency of the soil nutrients also had a significant impact on crop yields. Compared with soybean, peanut was more suitable for intercropping with apple trees to obtain economic benefits in the region. We concluded that apple-soybean and apple-peanut intercropping systems can be practical and beneficial in the region. However, the distance between crops and tree rows should be adjusted to minimize interspecies competition. Agronomic measures such as regular canopy pruning, root barriers, additional irrigation and fertilization also should be applied in the intercropping systems.     

Tree photosynthesis modulates soil respiration on a diurnal time scale

To estimate how tree photosynthesis modulates soil respiration, we simultaneously and continuously measured soil respiration and canopy photosynthesis over an oak‐grass savanna during the summer, when the annual grass between trees was dead. Soil respiration measured under a tree crown reflected the sum of rhizosphere respiration and heterotrophic respiration; soil respiration measured in an open area represented heterotrophic respiration. Soil respiration was measured using solid‐state CO₂ sensors buried in soils and the flux‐gradient method. Canopy photosynthesis was obtained from overstory and understory flux measurements using the eddy covariance method. We found that the diurnal pattern of soil respiration in the open was driven by soil temperature, while soil respiration under the tree was decoupled with soil temperature. Although soil moisture controlled the seasonal pattern of soil respiration, it did not influence the diurnal pattern of soil respiration. Soil respiration under the tree controlled by the root component was strongly correlated with tree photosynthesis, but with a time lag of 7–12 h. These results indicate that photosynthesis drives soil respiration in addition to soil temperature and moisture.     

Photosynthetic acclimation to simultaneous and interacting environmental stresses along natural light gradients: optimality and constraints

There is a strong natural light gradient from the top to the bottom in plant canopies and along gap-understorey continua. Leaf structure and photosynthetic capacities change close to proportionally along these gradients, leading to maximisation of whole canopy photosynthesis. However, other environmental factors also vary within the light gradients in a correlative manner. Specifically, the leaves exposed to higher irradiance suffer from more severe heat, water, and photoinhibition stresses. Research in tree canopies and across gap-understorey gradients demonstrates that plants have a large potential to acclimate to interacting environmental limitations. The optimum temperature for photosynthetic electron transport increases with increasing growth irradiance in the canopy, improving the resistance of photosynthetic apparatus to heat stress. Stomatal constraints on photosynthesis are also larger at higher irradiance because the leaves at greater evaporative demands regulate water use more efficiently. Furthermore, upper canopy leaves are more rigid and have lower leaf osmotic potentials to improve water extraction from drying soil. The current review highlights that such an array of complex interactions significantly modifies the potential and realized whole canopy photosynthetic productivity, but also that the interactive effects cannot be simply predicted as composites of additive partial environmental stresses. We hypothesize that plant photosynthetic capacities deviate from the theoretical optimum values because of the interacting stresses in plant canopies and evolutionary trade-offs between leaf- and canopy-level plastic adjustments in light capture and use.     

Importance of canopy structure on photosynthesis in single- and multi-species assemblages of marine macroalgae

Plant communities utilize available irradiance with different efficiency depending not only on their photosynthetic characteristics but also on the canopy structure and density. The importance of canopy structure are well studied in terrestrial plant communities but poorly studied in aquatic plant communities. The objective of this study was to evaluate macroalgal community photosynthesis in artificial constructed communities of one to four species with different morphologies along a range of leaf (i.e.=thallus) area densities. In a laboratory set-up we measured net photosynthesis and dark respiration in constructed assemblages of macroalgae, excluding effects other than photosynthesis of individual tissue and distribution of photons in the canopy from influencing metabolism. We hypothezised that 1) canopy structure determines the actual rates of photosynthesis relative to the optimal rates and 2) multi-species communities attain higher maximum photosynthetic rates than single species communities. We found that differences in canopy structure outweighed large differences in tissue photosynthesis resulting in relatively similar maximum community photosynthetic rates among the different single and multi-species assemblages (20.1–40.5 μmol O₂ m⁻² s⁻¹). Canopy structure influenced community photosynthesis both at low and high leaf area densities because it determines the ability of macroalgae to use the photosynthetic potential of their individual tissues. Due to an averaging effect the photosynthetic rate at high leaf area density was more similar among multi-species community than among single-species communities. Multi-species communities had, on average, a slightly higher photosynthetic production than expected from photosynthesis of single species communities. Moreover multi-species communities were capable of exposing new tissue to irradiance up to high densities thereby avoiding a decrease in net photosynthesis. This finding suggests that multi-species communities may be able to maintain higher biomass per unit ground area than single-species communities.     

The diel imprint of leaf metabolism on the δ13 C signal of soil respiration under control and drought conditions

Recent (13) CO(2) canopy pulse chase labeling studies revealed that photosynthesis influences the carbon isotopic composition of soil respired CO(2) (δ(13) C(SR)) even on a diel timescale. However, the driving mechanisms underlying these short-term responses remain unclear, in particular under drought conditions. The gas exchange of CO(2) isotopes of canopy and soil was monitored in drought/nondrought-stressed beech (Fagus sylvatica) saplings after (13) CO(2) canopy pulse labeling. A combined canopy/soil chamber system with gas-tight separated soil and canopy compartments was coupled to a laser spectrometer measuring mixing ratios and isotopic composition of CO(2) in air at high temporal resolution. The measured δ(13) C(SR) signal was then explained and substantiated by a mechanistic carbon allocation model. Leaf metabolism had a strong imprint on diel cycles in control plants, as a result of an alternating substrate supply switching between sugar and transient starch. By contrast, diel cycles in drought-stressed plants were determined by the relative contributions of autotrophic and heterotrophic respiration throughout the day. Drought reduced the speed of the link between photosynthesis and soil respiration by a factor of c. 2.5, depending on the photosynthetic rate. Drought slows the coupling between photosynthesis and soil respiration and alters the underlying mechanism causing diel variations of δ(13) C(SR).     

Acclimation of photosynthesis in canopies: models and limitations

Within a time-scale of several days photosynthesis can acclimate to light by variation in the capacity for photosynthesis with depth in a canopy or by variation in the stoichiometry of photosynthetic components at each position within the canopy. The changes in leaf photosynthetic capacity are usually related to and expressed as changes in leaf nitrogen content. However, photosynthetic capacity and leaf nitrogen never match exactly the photon flux density (PFD) gradient within a canopy. As a result, photosynthetic light use efficiency, i.e. photosynthetic performance per incident PFD, increases considerably from the top of the canopy to the lower shaded part. Many of existing optimisation models fail to express the actual pattern of nitrogen or photosynthetic capacity distribution within a canopy. This failure occurs because these optimisation models do not consider that the quantitative aspect of photosynthesis acclimation is a whole plant phenomenon. Although turnover models, which describe the distribution of the photosynthetic apparatus within a canopy as a dynamic equilibrium between breakdown and regeneration of apparatus with respect to nitrogen availability, photosynthetic rate and export of carbohydrates, produce realistic results, these models require confirmation. The mechanism responsible for changes in the relative share of light-harvesting apparatus as acclimation to irradiance remains unknown. Ability of the photosynthetic apparatus to balance properly the light harvesting capacity with electron transport and biochemical capacities is limited. As a result of this fundamental limitation, photosynthetic light use efficiency always increases with increasing thickness of the photosynthetic apparatus.     

不同土壤水分下山杏光合作用CO2响应过程及其模拟

在半干旱黄土丘陵区,以2年生盆栽山杏为材料,应用CIRAS-2型光合作用系统,测定了8个土壤水分梯度下山杏光合作用的CO₂响应过程,并采用直角双曲线模型、指数方程和直角双曲线修正模型对其CO₂响应数据进行拟合,分析了山杏光合作用与土壤水分的定量关系.结果表明: 山杏CO₂响应过程对土壤水分有明显的阈值响应特征.维持山杏叶片较高的光合速率(P_n)和羧化效率(CE)的土壤相对含水量(RWC)在46.3%～81.9%,在此水分范围内,光合作用没有发生明显的CO₂饱和抑制现象;当RWC超出此范围,土壤水分升高或降低均明显降低山杏叶片的光合能力(P_{n max})、CE和CO₂饱和点(CSP).在不同土壤水分条件下,3个模型对山杏CO₂响应数据的模拟效果有明显差别.在46.3%～81.9%土壤水分范围内,3个模型均能较好地拟合山杏CO₂响应过程及其特征参数CE、CO₂补偿点(Γ)和光呼吸速率(R_p),其拟合精度均表现为直角双曲线修正模型＞指数方程＞直角双曲线模型;当土壤水分含量过高(RWC＞81.9%)或过低(RWC＜46.3%)时,只有直角双曲线修正模型能较好地拟合山杏CO₂响应过程及其特征参数.RWC在46.3%～81.9%范围内,山杏具有较高的光合作用效率;与传统直角双曲线模型和指数方程相比,直角双曲线修正模型具有更好的适用性.     

Implications of shoot structure on the rate of photosynthesis at different levels in a coniferous canopy using a model incorporating grouping and penumbra

1. The connection between high leaf area index (LAI) and photosynthetic production with two attributes of coniferous canopy structure: small leaf size and grouping of needles on shoots, was analysed using a simulation model.
2. The small size of conifer needles gives rise to penumbras, which even out the distribution of direct sunlight on the leaf area and thereby act to increase the rate of canopy photosynthesis per unit of LAI.
3. Grouping, by producing a non-uniform distribution of leaf area, causes a decrease in total canopy light interception at any given LAI, but improves the photosynthetic light capture by shoots in the lower canopy.
4. Application of the model on a case study showed that: (a) grouping had a negative effect on the rate of photosynthesis in the upper canopy, but deeper down in the canopy the situation was reversed; (b) in the lower canopy, photosynthetic rates were up to 10 times higher as a result from the combined effect of grouping and penumbra; (c) grouping did not improve the rate of canopy photosynthesis per unit of LAI, however, it can have a positive effect on the total photosynthetic production by allowing a higher productive LAI to be maintained; (d) penumbra, on the other hand, increased the rate of canopy photosynthesis by as much as 40% for moderate values of the LAI.     

羊草(Leymus chinensis)叶片光合参数对干旱与复水的响应机理与模拟

利用典型草原优势植物羊草（Leymus chinensis）对不同水分胁迫与复水响应的植物光合生理生态模拟实验与野外观测资料,分析研究了羊草叶片光合参数Kcmax（Rubisco的最大羧化速率）、Jmax（最大光合电子传递速率）和TPU（磷酸丙糖利用率）对干旱与复水的响应机理。结果表明,无论是模拟实验还是野外观测均显示羊草叶片的光合参数随着土壤水分的增加呈抛物线曲线变化,但各光合参数最大值对土壤水分的响应不同。温室模拟下的羊草光合参数Vcmax,Jmax和TPU在土壤含水量分别在15．56％,15．89％和16．23％时达到最大,而野外观测羊草的光合参数Vcmax,Jmax和TPU在土壤含水量分别为16．89％,17％和16．79％时达到最大。复水后羊草植株叶片光合参数的变化取决于前期干旱的影响,土壤含水量18％～19％和15％～16％处理的羊草复水后光合参数能够恢复正常,前者甚至超过正常水平,说明适宜的水分胁迫在复水后能够提高羊草叶片的光合能力,促进光合作用;土壤含水量10％～12％和7％～9％处理下的羊草复水后光合参数则不能恢复到正常水平。土壤含水量15％～16％可能是羊草光合能力在水分胁迫后能否恢复的阈值。     

间伐改形对陇东旱塬密闭苹果园树体冠层结构和发育后期叶片质量的影响

为研究间伐改形对红富士成龄乔化密植果园树体冠层特征、生育后期叶片生理特性、养分积累分配规律、光合生产力和土壤水分时空分布动态的影响,以18年生‘红富士’苹果密闭园为试材,对苹果树单株结构参数、枝量、枝类组成、覆盖率、叶片光合速率等参数进行测定.结果表明: 密闭果园叶片光合作用受非气孔因素限制,导致PSII最大光化学效率、PSII光合潜能、光合性能指数下降了1.2%、11.5%、13.9%.间伐改形后,叶面积指数、树冠覆盖率有所降低,使得冠层直射光透过系数增加了79.0%,树形结构有所改善.苹果园总枝数降低到约1100400条·hm^-2,单株枝量增加了5.0%,短枝比例提高至73.0%.由于冠层光照条件的改善,叶面积、比叶质量、百叶重、叶绿素含量有不同程度的提高.叶片光合速率的提高促进了光合产物的积累,淀粉、蛋白质含量为密闭果园的143.5%、107.8%.叶片的发育质量与其所处的光照辐射环境有着密切联系,密闭果园经间伐、改形后,果园群体结构和冠层光照得以改善,促进了叶片生长发育,提高了叶片光合效能,降低了果园土壤水分的无效消耗,是陇东旱塬苹果产区密闭果园适宜的调整、优化方案.